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Alan Dugatkin
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Chapter 01: An Overview of Evolutionary Biology
MULTIPLE CHOICE
1. Charles Darwin’s theory of evolution resulted in a because it changed the way we

understand how all living things came to be and how they function.
a. paradigm shift c. physical theory
b. null hypothesis d. static discovery
ANS: A DIF: Moderate REF: 1.0
OBJ: 1.0.a. Explain the paradigm shift that occurred in biology when Darwin laid out his theory of
evolution. MSC: Remembering
2. Darwin realized that the often exquisite fit of species to their environment is primarily the result of
a. paradigm shifts. c. supernatural events.
b. natural selection. d. empirical research.
ANS: B DIF: Easy REF: 1.0
3. In describing the importance of Darwin’s theory of evolution, Theodosius Dobzhansky argued that
a. nothing in biology makes sense except in the light of evolution.
b. this theory of evolution changes the way we think of Earth in the universe.
c. evolution can now be used to control our destiny.
d. we no longer need to invoke the supernatural for any cause.
4. Darwin’s theory of evolution provided explanations for all of the following EXCEPT
a. the diversity of life on the planet.
b. why the vast majority of species are extinct.
c. why Earth is not the center of the universe.
d. the similarities and differences among species.
ANS: C DIF: Moderate REF: 1.0
evolution. MSC: Understanding
5. What is evolutionary biology?

a. a description of the fossil record
b. the field of science that deals with matter and its motion through space and time
c. the study of interactions among organisms and their environment
d. the study of the origin, maintenance, and diversity of life
ANS: D DIF: Easy REF: 1.1
OBJ: 1.1.a. Describe what evolutionary biology is. MSC: Remembering
6. To understand the evolution of a species, we need to know about its ancestors and what changes have
occurred along the way. Darwin called this process
a. natural selection. c. genetics.
b. descent with modification. d. changes in fitness.
ANS: B DIF: Moderate REF: 1.1
OBJ: 1.1.a. Describe what evolutionary biology is. MSC: Remembering
7. Which of the following best summarizes the process of evolution by natural selection?
a. Most mutations have harmful effects and therefore increase in frequency over time.
b. Mutations that do not affect the fitness of individuals are selected to decrease in frequency
over time.
c. Natural selection results in mutations that disrupt finely tuned processes.
d. Mutations that improve the fitness of individuals will tend to increase in frequency over
time.
ANS: D DIF: Moderate REF: 1.1
OBJ: 1.1.a. Describe what evolutionary biology is. MSC: Understanding
8. Which process is demonstrated in the figure shown?
a. artificial selection c. genetics

b. natural selection d. paradigm shift
OBJ: 1.1.b. Compare artificial selection with natural selection in regard to the domestication of crops
and livestock. MSC: Understanding
9. In what way are natural selection and artificial selection similar?

a. Both result in increased frequency of beneficial traits.
b. Both rely on human intervention to prevent deleterious mutations from increasing in
frequency.
c. Both have been occurring for the past 3.5 billion years.
d. Neither are able to produce dramatic changes in traits.
and livestock. MSC: Analyzing
10. In what way is artificial selection different from natural selection?

a. Artificial selection cannot result in changes in traits.
b. Artificial selection relies on humans choosing which traits are beneficial.
c. Artificial selection is only applied to plants.
d. Artificial selection results in organisms that cannot survive and reproduce.
11. Which of the following is an example of artificial selection?

a. selection for pests resistant to pesticides
b. selection for weeds that can grow in the presence of herbicides applied by farmers
c. selection for cows that produce more milk
d. selection for insects that transmit disease more efficiently
and livestock. MSC: Applying
12. Looking at the figure, predict what will happen if a new antibiotic to the food poisoning bacteria
Campylobacter jejuni is developed and prescribed heavily by physicians, beginning this year.
a. Campylobacter jejuni will be effectively treated for the next two to three decades.
b. Campylobacter jejuni will be effectively treated for the next two to five years.
c. The new antibiotic will make all former antibiotics for treating Campylobacter jejuni
obsolete.
d. The new antibiotic will never work to treat Campylobacter jejuni.
OBJ: 1.1.c. Explain how evolutionary biology informs our understanding of antibiotic resistance.
MSC: Applying
13. Antibiotic resistance evolves quickly because

a. antibiotics impose strong selection for resistance.
b. antibiotics cause new mutations in bacteria that result in antibiotic resistance.
c. natural selection causes evolution to slow down.
d. bacterial infections should not be treated with antibiotics.
MSC: Understanding
14. Why do physicians often prescribe antibiotics in combination?

a. to decrease the rate at which antibiotic resistance evolves and spreads
b. to prevent common side effects to the patient
c. to increase the likelihood that resistance mutations will arise in the bacteria
d. in case an infection is caused by both bacteria and viruses
MSC: Understanding
15. A branching tree showing historical relationships among species is known as a(n)
a. species network. c. phylogenetic tree.
b. natural selection tree. d. extinction chart.
ANS: C DIF: Easy REF: 1.1
OBJ: 1.1.d. Explain how a phylogenetic perspective can change how scientists address the challenges
of conservation biology. MSC: Remembering
16.
Each tip of the phylogenetic tree shown represents
a. extinction. c. living or extant taxon.
b. domain. d. speciation event.
of conservation biology. MSC: Remembering
17. According to the phylogenetic tree shown, fungi are most closely related to which of the following?
a. plants c. animals
b. amoebas d. bacteria
of conservation biology. MSC: Applying
18. If you could protect from extinction only the lineages derived from two of the nodes in the figure,
which pair would you save to yield the greatest phylogenetic diversity?
a. Nodes E and F c. Nodes C and D
b. Nodes D and E d. All of the choices are equivalent.
19. If we are interested in conserving phylogenetic diversity, the extinction of which group of species in
the figure shown is a greater loss?
a. B and I c. G and J
b. E, F, and I d. B and C
20. The loss of which species on the figure would represent the loss of the most significant phylogenetic
diversity?
a. A c. C
b. B d. D
ANS: A DIF: Difficult REF: 1.1
21. What are the two major types of empirical research in evolutionary biology?
a. mutation and natural selection c. empirical and theoretical
b. observation and manipulation d. experiments and research
ANS: B DIF: Easy REF: 1.2
OBJ: 1.2.a. Define “empirical research” and describe its two main categories.
MSC: Remembering
22. Which of the following is an example of observational research?

a. removing all of the bee pollinators from a field and watching what happens to the other
pollinators
b. calculating the predicted effect of different species of pollinators
c. measuring the behavior of different pollinators visiting flowers in a field
d. adding new flowers to a field and recording the effect on pollinators
MSC: Understanding
23. Manipulative experiments

a. allow scientists to directly assess how changes in one component of a system influence the
other components.
b. allow scientists to examine only correlations among data.
c. require altering multiple variables at the same time.
d. must be performed under laboratory conditions.
MSC: Understanding
24. More than 100 years ago, Charles Darwin and his colleague Thomas Huxley hypothesized that humans
share a common ancestor, based on
a. anatomical evidence.
b. fossil evidence gathered during Darwin’s voyage on the HMS Beagle.
c. manipulative experiments.
d. mathematical modeling.
ANS: A DIF: Easy REF: 1.2
OBJ: 1.2.b. Explain how molecular genetics informs our understanding of the relationships among
humans and great apes. MSC: Remembering
25. Humans have 23 pairs of chromosomes and chimps have 24 pairs. How has this difference best been
explained?
a. Humans and chimps are not related.
b. Chimps gained a chromosome during their evolution, which explains why they differ from
humans.
c. Humans and chimps share very little genetic material.
d. Humans have one chromosome that is the result of a fusion of two chromosomes in our
ancestors.
humans and great apes. MSC: Remembering
26. If chimpanzee and human genomes differ only about 1.3% at the level of DNA base pairs, how might
we explain the dramatic differences in appearance, behavior, cultures, and so on between humans and
chimps?
a. There is a correlation between increased rates of divergence with known functions of
alleles in humans and chimps.
b. Natural selection has been acting on clusters of genes associated with both survival and
reproduction in humans and chimps.
c. Important differences exist in the expression of genes in humans and chimps.
d. Humans and chimps have been diverging from each other for much longer than they have
with other primates.
humans and great apes. MSC: Understanding
27. If testes size is correlated with number of sperm produced, under what condition is larger testes size
more likely to be evolutionarily advantageous?
a. Females mate with a single male. c. Females mate with multiple males.
b. Males mate with a single female. d. Males mate with multiple females.
OBJ: 1.2.c. Describe how primate breeding systems can influence the evolution of testes size.
MSC: Understanding
28. In the figure below, the lighter gray circles represent single-male breeding systems in primates and the
darker gray circles indicate multi-male systems. Which of the following statements is consistent with
these data?
a. Single-male breeding systems select for smaller testes size because sperm are not needed
to increase reproductive success.
b. Body weight does not influence testes size because selection only favors large testes in
multi-male systems.
c. Species with multi-male systems evolve larger testes because they have higher body
weights.
d. Selection favors large testes relative to body weight when a male’s sperm must compete
directly with other males.
ANS: D DIF: Difficult REF: 1.2
OBJ: 1.2.c. Describe how primate breeding systems can influence the evolution of testes size.
MSC: Analyzing
29. Mathematical models are useful for

a. disproving the theory of evolution by natural selection.
b. providing observations of the natural world.
c. manipulating experimental conditions in the laboratory.
d. helping us understand how complicated systems work.
ANS: D DIF: Easy REF: 1.2
OBJ: 1.2.d. Describe the role of mathematical models of biological systems in evolutionary biology.
MSC: Understanding
30. In which of the following scenarios are models being used to make predictions and plan for the future?
a. A policeman clocks the speed of a motorist with a radar gun.
b. Someone checks the weather forecast for the weekend.
c. An evolutionary biologist estimates fitness by measuring gene frequencies over time.
d. A scientist changes the number of males and females in a population and observes the
evolutionary response.
MSC: Applying
31. Sir Ronald A. Fisher developed a sex ratio model partly because so much observational data suggested
that the 1:1 sex ratio was common in nature and he wanted to understand why. What is the natural
ordering when it comes to empirical and theoretical approaches?
a. Good theory should postdate data collecting.
b. Good theory should precede data collecting.
c. Good theory can either precede or postdate data collecting.
d. Good theory should precede hypothesis collecting.
MSC: Understanding
32. In developing his sex ratio theory, Sir Ronald A. Fisher assumed that sex ratio is under genetic control.
Why is this an important assumption?
a. Only traits under genetic control can evolve by natural selection.
b. Sex ratio is under genetic control in humans.
c. Fisher wanted to include every possible variable in his model.
d. Traits that are not under genetic control cannot affect an individual’s reproductive success.
OBJ: 1.2.e. Explain why many species display an even sex ratio.
MSC: Understanding
33. Imagine a population that has 50 males and 25 females. Which of the following parental sex ratio
strategies will be most successful?
a. produce all sons
b. produce all daughters
c. produce half sons and half daughters
d. produce 2/3 sons and 1/3 daughters
MSC: Applying
34. Sir Ronald A. Fisher’s prediction that sex ratios should be 1:1 relies on the principle that
a. the genetic basis of sex is chromosomal.
b. total reproductive success of males is higher than that of females.
c. male births are less common than female births.
d. the rarer sex will have better mating prospects than the more common sex.
MSC: Understanding
35. During the observations of a species of blue moon butterflies on the Samoan island of Upolu, 99% of
the butterflies were female and only 1% were male. Only five years later the male:female sex ratio was
1:1. What caused this?
a. Wolbachia lost the ability to kill all male butterflies.
b. Genetic changes in the Upolu butterflies resulted in suppression of the male-killing effect
of Wolbachia.
c. Upolu butterflies evolved to become resistant to Wolbachia infection.
d. Upolu was recolonized by butterflies from the nearby island Savali, where sex ratios were
closer to 1:1.
MSC: Understanding
SHORT ANSWER
1. In On the Origin of Species, Charles Darwin brought together two ideas that resulted in a paradigm
shift in biology. What were these two ideas?
ANS:
1. The diversity of life has descended from preexisting species, which share a common ancestor.
2.The fit of species to their environments is primarily the result of natural selection.
DIF: Easy REF: 1.0

2. Describe Charles Darwin’s theory of evolution by natural selection.
ANS:
A gradual process in which forms of organisms that are better suited to their environment increase in
frequency in a population over sufficient periods of time.
DIF: Easy REF: 1.0

3. Charles Darwin’s theory of natural selection is considered a paradigm shift—a theory that has
wide-ranging effects. Describe another paradigm shift that has occurred in biology. Make a case for
why this shift has fundamentally changed the way scientists see the world and the sorts of questions
they ask.
ANS:
Answers will vary. An example from the text includes the following: when astronomers discovered
that Earth was not at the center of the universe, the way that we thought of Earth and our place in
nature was radically changed.
DIF: Moderate REF: 1.0

evolution. MSC: Applying
4. Sources for the data that evolutionary biologists use to test their hypotheses are derived from many
subdisciplines of the biological sciences. Identify the five data sources shown in the photographs.
ANS:
A. Data from the fossil record (Paleontology); B. behavioral data; C. morphological data
(Morphology); D. embryological data (Embryology); and E. molecular data (Genetics).
DIF: Moderate REF: 1.1 OBJ: 1.1.a. Describe what evolutionary biology is.
MSC: Remembering
5. How has artificial selection been used to shape the characteristics of food-producing plants?
ANS:
Using the most desirable plants as parental stock for the next generation has resulted in plants that are,
for example, hardier, quicker growing, and better tasting.
DIF: Easy REF: 1.1

and livestock. MSC: Applying
6. Charles Darwin used the process of artificial selection during domestication as an analogy to explain
natural selection. In what ways are artificial and natural selection similar? In what ways do they differ?
ANS:
They are similar in that both processes take the best individuals to be parents for the next generation
and thus the most desirable traits will increase in frequency. They differ in what makes individuals the
“best” and what traits are “desirable.” Under natural selection, the best individuals are those that have
the highest survival and reproductive rates. Under artificial selection, the breeder decides what
individuals and traits are best.

7. Given the fact that humans are the ones producing and distributing pesticides, why do we call the
development of pesticide resistance natural selection rather than artificial selection?
ANS:
The distinction between natural and artificial selection refers not to whether humans are involved but
to whether they deliberately choose which organisms will survive and reproduce.

and livestock. MSC: Understanding
8. The majority of antibiotic use in the United States is for agriculture, primarily in livestock. Why is this
a problem?
ANS:
Antibiotic-resistant strains of bacteria can evolve in farm animals and then spread to humans,
decreasing the probability of having an effective antibiotic at hand for use in human infections.

MSC: Understanding
9. If you are interested in preserving phylogenetic diversity as part of a conservation strategy, would you
be more concerned about the loss of all of the species labeled with an X or of the one species labeled
with a Y?
ANS:
Loss of the Y species would produce a greater loss of phylogenetic diversity than loss of all of the X
species.

10. Evolutionary biologists, like most scientists, use two empirical approaches to study evolution,
observation and manipulation. What is the difference between these approaches?
ANS:
Observation involves gathering data without making any changes in a natural system. Manipulation
requires changing one or more components of a system to determine its/their effects.
DIF: Easy REF: 1.2

MSC: Understanding
11. In performing manipulative experiments, scientists usually try to alter only one variable at a time. Why
do they do this?
ANS:
By changing only one variable at a time and observing the consequences of that change, the researcher
is better able to determine the specific effect of each variable.
DIF: Easy REF: 1.2

MSC: Understanding
12. In comparing pairs of genes in the human and chimp genomes, Tarjei Mikkelsen and his colleagues
first determined the expected degree of divergence between the two genomes based on the
accumulation of neutral mutations. Why is this an important first step in understanding how evolution
is occurring in these species?
ANS:
The rate of neutral evolution serves as a baseline for comparison. If some pairs of genes are evolving
faster than this baseline rate, then scientists could infer that those genes are not evolving neutrally but
rather have experienced natural selection.

13. A comparison of gene clusters in humans and chimps from Tarjei Mikkelsen and colleagues revealed
that some types of genes were evolving particularly rapidly. Name one type of gene cluster that they
found to evolve rapidly and explain why you think it shows this type of evolution.
ANS:
Possible answers include the following: Gene clusters associated with survival and reproduction
evolve rapidly, likely because selection is strongest on genes directly affecting fitness. Gene clusters
associated with disease resistance are evolving rapidly, likely due to the constantly changing selective
pressure from coevolving parasites and pathogens.
DIF: Difficult REF: 1.2

14. Sir Ronald A. Fisher’s sex ratio model predicts a 1:1 female:male sex ratio. Under what conditions
might you expect a different ratio to evolve?
ANS:
If one of the assumptions of Fisher’s model is not met in a system, then the ratio may not be 1:1. For
example, if the cost of producing and raising males differs from that of females, then we might expect
a different equilibrium sex ratio.

MSC: Understanding
15. In a population with 50 males and 25 females that produce a total of 100 offspring, calculate how
many grandoffspring per child a parent will have if (a) the parent produces half sons and half
daughters, or (b) the parent produces 1/4 sons and 3/4 daughters. Which strategy will be favored by
natural selection in this population?
ANS:
Using the equation k(Nm) + (1−k)(Nf), we calculate that parent (a) will produce:
(12)(10050) + (12)(10025) = 3 grandoffspring per child
and parent (b) will produce:
(14)(10050) + (34)(10025) = 3.5 grandoffspring per child.
Parent (b) will be favored because it produces more offspring of the rare sex.

MSC: Applying
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ornament; but the main figures and general design have perished.
The walls of the opposite chamber were never cased with marble, so
that the pilgrims were able to leave here the same tokens of their
visits as they left at St. Sixtus’. The graffiti are of the same general
character, but of a somewhat later date; the old forms of prayer have
disappeared; most of the names and inscriptions are in Latin; and
among the few that are Greek, there are symptoms of Byzantine
peculiarities.
The chief object of interest, however, now remaining in these
chambers is the epitaph which stands in the middle of the smaller
room. Of course, this was not its original position; but it has been so
placed, in order that we may see both sides of the stone without
difficulty, for both are inscribed. The stone was originally used for an
inscription in honour of Caracalla, belonging to the year 214. The
Christian inscription on the other side professes to have been set up
by “Damasus, Bishop, to Eusebius, Bishop and Martyr,” and to have
been written by Furius Dionysius Filocalus, “a worshipper (cultor)
and lover of Pope Damasus.” But it is easy to see at a glance that it
never was really executed by the same hand to which we are
indebted for so many other beautiful productions of that Pope. At
first, therefore, and whilst only a few fragments of this inscription had
been recovered, De Rossi was tempted to conjecture that it might be
one of the earliest efforts of the artist who subsequently attained
such perfection. At length, however, the difficulty was solved in a
more sure and satisfactory way. A diligent search in the earth with
which the chamber was filled brought to light several fragments of
the original stone, on which the letters are executed with the same
faultlessness as on the other specimens of its class. The visitor to
the Catacombs may see them painted, in a different colour from the
rest, in the copy of the epitaph which De Rossi has caused to be
affixed to the wall; and he will observe that amongst them are some
letters which are wanting in the more ancient copy transcribed on the
reverse of Caracalla’s monument. It is clear that the original must
have been broken in pieces, by the Lombards or other ancient
plunderers of the Catacombs, and that the copy which we now see is
one of the restorations by Pope Vigilus or some other Pontiff about
that time (page 47). The copyist was so ignorant that he could only
transcribe the letters which were on the spot before his eyes, and,
even when he was conscious that a letter was missing, he could only
leave a vacant space, being doubtful how it should be supplied.
Witness the space left for the first letter of Domino in the penultimate
line of the inscription, and the word in altogether omitted in the third
line.
“Heraclius forbad those who had fallen away [in times of

persecution] to grieve for their sins.
But Eusebius taught those unhappy men to weep for their crimes.
The people are divided into parties; fury increases;
Sedition, murder, fighting, quarrelling, and strife.
Presently both [the Pope and the heretic] are exiled by the cruelty
of the tyrant,
Although the Pope was preserving the bonds of peace inviolate.
He bore his exile with joy, looking to the Lord as his Judge.
And on the shore of Sicily gave up the world and his life.”
Having sufficiently considered the form of the inscription, let us

now say a few words about its substance, which is important,
because it restores to us a lost chapter of Church history. Every
student knows how keenly contested in the early ages of the Church
was the question as to the discipline to be observed towards those
Christians who relapsed into an outward profession of Paganism
under the pressure of persecution. There were some who would fain
close the door of reconciliation altogether against these unhappy
men (miseri), whilst others claimed for them restitution of all
Christian privileges before they had brought forth worthy fruits of
penance.
The question arose whenever a persecution followed after a long
term of peace; for during such a time men’s minds were specially apt
to decline from primitive fervour, and the number of the lapsed to
increase. We are not surprised, therefore, to find the question
agitated during the persecution of Decius in the middle of the third
century. There is still extant a touching letter, written to St. Cyprian
by the clergy of Rome at a time when the Holy See was vacant after
the martyrdom of St. Fabian, which clearly defines the tradition and
practice of the Church. In it they say that absolution was freely given
to those of the lapsed who are in danger of death, but to others only
when wholesome penance has been exacted; and they declare that
“they have left nothing undone that the perverse may not boast of
their being too easy, nor the true penitents accuse them of inflexible
cruelty.” The same question arose under the same circumstances in
the persecution of Diocletian. Pope Marcellus was firm in upholding
the Church’s discipline, but he was resisted with such violence that
public order was disturbed in the city by the strife of contending
factions, and the Pope was banished by order of the Emperor
Maxentius. This we learn from another inscription of Pope Damasus,
who says that he wrote it in order that the faithful might not be
ignorant of the merit of the holy Pontiff. Eusebius was the immediate
successor of Marcellus, and the epitaph now before us is clearly a
continuation of the same history, ending in the same punishment of
the Pope, as the reward of his contention for the liberties of the
Church. For it should be remembered that these Popes were driven
from their see and died in exile, not because they refused to
apostatize, but because they insisted on maintaining the integrity of
ecclesiastical discipline. They may justly be reckoned, therefore,
among the earliest of that noble army of martyrs, who, from those
days even to our own, have braved every danger rather than
consent to govern the Church in accordance with other than the
Church’s rules.
It yet remains to make two further remarks upon the epitaph of
Pope Eusebius before we leave it. The first is, that he is called a
martyr, though it nowhere appears that he really shed his blood; but
this is by no means the only instance in which the title of martyr is
given in ancient documents to men who have suffered for the faith
and died whilst those sufferings continued. And secondly, it is to be
observed that although we have no record of the translation of the
body of St. Eusebius from Sicily to Rome, there is no reason to
doubt the fact. All the earliest monuments speak of him as buried in
a crypt of the Cemetery of St. Callixtus, and although the law forbad
the translation of the bodies of those who had died in exile unless
the emperor’s permission had been previously obtained, the old
lawyers tell us that this permission was freely given. Numerous
examples teach us the great anxiety of the ancient churches to have
their bishops buried in the midst of them; no doubt, therefore, the
necessary permission was asked for, as soon as a change in the
imperial policy towards the Church made it possible; and the body of
St. Eusebius was recovered and brought to Rome soon after his
death, just as that of one of his predecessors, St. Pontian, had been
brought from Sardinia by St. Fabian.
CHAPTER IV.
THE TOMB OF ST. CORNELIUS.
We have not promised to conduct the visitor to everything that is

worth seeing in this cemetery, but only to enumerate and explain the
principal monuments of historical importance which every stranger
usually sees. And the only specimen of this class which remains to
be spoken of is the tomb of St. Cornelius, which lies some way off. In
order to reach it we must traverse a vast network of galleries, narrow
and irregular, connecting what were once independent cemeteries,
or at least were areæ added at various times to the Cemetery of
Callixtus. If our guide is not in too great haste, he may allow us to
step aside into two or three chambers by the way, in which are
certain objects of interest worth looking at. The first is a long
inscription belonging to the last decade of the third century, in which
the Deacon Severus records that he has obtained leave from the
Pope Marcellinus to make a double chamber, with arcosolia and a
luminare, in which himself and his family may have quiet graves
(mansionem in pace quietam). This is in the third area of the
cemetery, next to the area in which we visited the crypt of St.
Eusebius.
In the adjoining area, and belonging probably to the same date, is
a very curious fresco, much damaged by having been cut through for
the sake of making a grave behind it, yet still easily distinguishable in
all its main features. The Good Shepherd occupies the centre of the
painting. On either side is an apostle, probably SS. Peter and Paul,
hastening away from Christ, Who has sent them to go and teach all
nations. These are represented by two sheep standing before each
of the apostles; and over their heads hangs a rock, whence pour
down streams of water, which the apostles are receiving in their
hands and turning on the heads of the sheep. We need no special
explanation of this; we have already learnt that the Rock is Christ,
and that the waters represent all Christian graces and sacraments.
But what is worth noticing in this picture is the various attitudes of the
sheep, and the corresponding distribution of the water. A perfect
torrent is falling on the animal that stands with outstretched neck and
head uplifted, drinking in all he hears with simplicity and eagerness;
whilst another, which has turned its back upon the apostle, is left
without any water at all. Of the other two, one is standing with head
downcast, as if in doubt and perplexity, and upon him too grace is
still being poured out more abundantly than upon the fourth, which is
eating grass, i.e., occupied with the affairs of this world.
On the right hand side of this arcosolium are two representations
of Moses; in the one he is striking the rock, and one of the Jews is
catching some of the water which gushes forth; in the other he is
taking off his shoes, preparing to obey the summons of God, who is
represented by a hand coming forth from the cloud. The painting on
the other side of the arcosolium is even more defaced than that in
the centre. A large semi-circular recess has been cut through it, and
then the smoke of the lamp which burnt in this recess during the
fourth and fifth centuries has almost obliterated the little that
remained of the figure of our Lord. He stood between two of His
apostles, who are offering Him bread and fish, and six baskets of
loaves stand on the ground before them.
And now we will not linger any more upon the road, but follow our
guide, who hurries forward along the intricate passages until he
lands us at last in an irregularly shaped space, illuminated by a
luminare, decorated with paintings, and bearing manifest tokens of
having been once a great centre of devotion. There is the pillar to
support the usual vessel of oil or more precious unguents to be burnt
before the tomb of the martyr; and hard by is a gravestone let into
the wall with the words Cornelius Martyr, Ep.
The stone does not close one of the common graves such as are
seen in the walls of the galleries or of the cubicula, neither is the
grave an ordinary arcosolium. The lower part of it, indeed, resembles
an arcosolium inasmuch as it is large enough to contain three or four
bodies, but there is no arch over it. The opening is rectangular, not
circular, and yet there is no trace of any slab having been let into the
wall to cover the top of the grave. It is probable, therefore, that a
sarcophagus once filled the vacant space, and that the top of this
sarcophagus served as the mensa or altar, an arrangement of which
other examples have been found.
But how came Pope Cornelius to be buried here, and not with his
predecessors in the Papal Crypt? He was Pope, a.d. 250, between
Fabian and Lucius, both of whom were buried, as we have seen, in
that crypt. It is to be observed, however, that Cornelius is the only
Pope, during the first three centuries, who bore the name of a noble
Roman family; and many ancient epitaphs have been found in the
area round this tomb, of persons who belonged to the same family. It
is obvious, therefore, to conjecture that this sepulchre was the
private property of some branch of the Gens Cornelia. The public
Cemetery of St. Callixtus may have been closed at this time by order
of the Government; but even without such a reason, it may have
been the wish of the family that the Pope should not be separated in
burial from the rest of his race. The same circumstance would
account for the epitaph being written in Latin, not in Greek, for many
of the old patrician families clung to the language of their forefathers
long after the use of Greek had come into fashion; and this departure
from the official language of the Church (for such, in fact, Greek
really was at that time) is quite of a piece with the preference of the
domestic to the official burial-place.
But whatever may be the true explanation of these circumstances,
the fact is at least certain that Cornelius was buried here; and above
and below the opening of his tomb are fragments, still adhering to
the wall, of large slabs of marble, containing a few letters of what
were once important inscriptions. The upper inscription was
unquestionably the work of Damasus. The letters of the lower,
though closely resembling the Damasine type, yet present a few
points of difference—sufficient to warrant the conjecture of De Rossi
that they were executed by the same hand, but with slight variations,
in order to mark that it belonged to another series of monuments. We
subjoin a copy of both inscriptions, in the form in which De Rossi
believes them to have been originally written. In the first inscription
the difference of type will distinguish the earlier half of each line,
which is a conjectural restoration, from the latter half which still
remains in situ; and in estimating the degree of probability of the
restorations, the reader should bear in mind two things: first, that the
Damasine inscriptions were engraved with such mathematical
precision that no emendations are admissible which would materially
increase or diminish the number of letters in each line; and secondly,
that whereas Damasus was in the habit of repeating himself very
frequently in his epitaphs, several of De Rossi’s restorations are
mere literal reproductions of some of his favourite forms of speech.
Had the following epitaph been found in some ancient MS., and
there attributed to Pope Damasus, we are confident that no critic
would have seen reason to doubt its genuineness:—
ASPICE, DESCENSU EXSTRUCTO TENEBRISQUE FUGATIS,

CORNELI MONUMENTA VIDES TUMULUMQUE SACRATUM.
HOC OPUS ÆGROTI DAMASI PRÆSTANTIA FECIT,
ESSET UT ACCESSUS MELIOR, POPULISQUE PARATUM
AUXILIUM SANCTI, ET VALEAS SI FUNDERE PURO
CORDE PRECES, DAMASUS MELIOR CONSURGERE
POSSET,
QUEM NON LUCIS AMOR, TENUIT MAGE CURA LABORIS.
“Behold, a new staircase having been made, and the darkness

put to flight,
You see the monuments of Cornelius and his sacred tomb.
This work the zeal of Damasus has accomplished, at a time when
he was sick;
That so the means of approach might be better, and the aid of the
saint
Put more within the reach of the people; and that if you pour forth
prayers
From a pure heart, Damasus may rise up in better health;
Though it has not been love of life, but rather anxiety for work,
that has retained him in this life.”
The second inscription De Rossi would restore as follows:—

SIRICIUS PERFECIT OPUS,
CONCLUSIT ET ARCAM
MARMORE, CORNELI QUONIAM
PIA MEMBRA RETENTAT
—that is to say, he supposes that, Damasus having died, his
successor Siricius completed the work that had been begun, and,
furthermore, strengthened the wall which enclosed the tomb of St.
Cornelius with this very thick slab of marble—a work which may
have been rendered necessary by the alterations already made by
Damasus. Of course, these restorations of the mutilated inscriptions
must always remain more or less doubtful, for we fear there is no
chance of any other fragments of the original ever coming to light.
We publish them under the same reserve with which he himself
proposes them, as at least approximations to the truth. He says that,
without daring to affirm their literal correctness, there are certainly
strong reasons for believing that they exactly reproduce the sense of
the original.
This same tomb of St. Cornelius will supply us with an example of
De Rossi’s power of happy conjecture, confirmed with absolute
certainty by subsequent discoveries. He had often publicly
expressed his confident expectation of finding at this tomb of St.
Cornelius some memorial of his cotemporary, St. Cyprian. These two
saints were martyred on the same day, though in different years; and
their feasts were, therefore, always celebrated together, just as they
are now, on the 16th of September, all the liturgical prayers for the
day being common to both. Now, De Rossi had found in one of the
old Itineraries, to whose accuracy of detail he had been greatly
indebted, an extraordinary misstatement, viz., that the bodies of both
these saints rested together in the same catacomb, whereas
everybody knows that St. Cyprian was buried in Africa. He
conjectured, therefore, that the pilgrim had been led into this blunder
by something he had seen at the tomb of St. Cornelius. On its
rediscovery, the cause of the error stands at once revealed.
Immediately on the right hand side of the grave are two large figures
of bishops painted on the wall, with a legend by the side of each,
declaring them to be St. Cornelius and St. Cyprian.
On the other side of the tomb is another painting, executed in the
same style, on the wall at the end of the gallery: two figures of
bishops, again designated by their proper names and titles. Only one
of these can now be deciphered, s̅c̅s̅ xustus p̅p̅ r̅o̅m̅, i.e., Pope
Sixtus II., of whose connection with this cemetery we have already
heard so often. The other name began with an O, and was probably
St. Optatus, an African bishop and martyr, whose body had been
brought to Rome and buried in this cemetery.
These paintings are manifestly a late work: perhaps they were
executed in the days of Leo III., a.d. 795-815, of whom it is recorded
in the Liber Pontificalis, that “he renewed the Cemetery of Sts. Sixtus
and Cornelius on the Appian Way;” and the legend which runs round
them would have a special significance as the motto of one who had
been almost miraculously delivered out of the hands of his enemies
by the Emperor Charlemagne. It is taken from the 17th verse of the
58th Psalm: “Ego autem cantabo virtutem Tuam et exaltabo
misericordiam Tuam quia factus es et susceptor meus.”... “I will sing
Thy strength, and will extol Thy mercy, for Thou art become my
support.” Of course, this had not been the earliest ornamentation of
these walls. Even now, we can detect traces of a more ancient
painting, and of graffiti upon it, underlying this later work. The graffiti
are only the names of priests and deacons, who either came here to
offer the holy sacrifice, or perhaps to take part in the translation of
the relics: “Leo prb., Theodorus prb., Kiprianus Diaconus,” &c.
We are drawing very near to the end of our subterranean walk:
indeed, the staircase which is to restore us to the upper air close to
the very entrance of the vineyard is immediately behind us, as we
stand contemplating the tomb of St. Cornelius. Nevertheless, if we
are not too weary, nor our guide too impatient, we should do well to
resist the temptation to escape, until we have first visited two small
chambers which are in the immediate neighbourhood. They contain
some of the most ancient specimens of painting to be found in the
whole range of the Catacombs. The ceilings are divided into circles
and other geometrical figures, and then the spaces are filled up with
graceful arabesques, birds, and flowers, peacocks, and dancing
genii. It was the sight of such paintings as these which led the
Protestant writer quoted in a former chapter to express an opinion
that, on first entering some of the decorated chambers in the
Catacombs, it is not easy to determine whether the work is Christian
or Pagan. Here, indeed, the Good Shepherd in one centre and
Daniel between two lions in the other soon solve the doubt; but all
the other details and the excellence of their execution may well have
suggested it. No one can doubt that the paintings belong to the very
earliest period of Christian art, when the forms and traditions of the
classical age had not yet died away.
In the first of the two chambers we are speaking of, there is
nothing special to be seen besides the ceiling; but the second and
more distant is more richly decorated. Here, two sepulchral
chambers open one into the other: over the doorway which admits to
the inner vault is represented the Baptism of our Lord by St. John:
He is coming up out of the water and the dove is descending upon
Him. On the wall opposite to the entrance is that fish carrying the
basket of bread and wine that has been already described (page 81).
On the wall to the left is a pail of milk standing on a kind of altar
between two sheep, and we know from St. Irenæus and from some
of the earliest and most authentic acts of the martyrs that milk was
an accepted symbol of the Holy Eucharist. Opposite to this are
doves and trees, which are often used as types of the souls of the
blessed in Paradise. Thus, on one side we have the faithful on earth
standing around the Divine food which prepares for heaven; and on
the other, souls released from the prison of the body have flown
away and are at rest, reposing amid the joys of another world; so
that it would almost seem as though the same sequence of ideas
presided over the decoration of these chambers, as was certainly
present to the minds of those who designed the ornamentation of the
sacramental chambers in the Cemetery of St. Callixtus (page 84).
And now at length we must conclude our visit to St. Callixtus. We
fear that we have already enumerated more than can be seen with
advantage during the course of a single visit; yet it is worth an effort
to see it all, because it includes monuments which illustrate nearly
every century of the period during which the Catacombs were used.
It is for this reason that a visit to St. Callixtus is so singularly
valuable, whether it be intended to take this cemetery as a sample of
all, or only to use it as an introduction to others. Those who propose
to pursue the subject further would do well to visit next the Catacomb
of SS. Nereus and Achilles, which lies at no great distance, off the
Via Ardeatina; then the Cemetery of Pretextatus on the other side of
the Via Appia; and finally, the Cœmeterium Ostrianum on the Via
Nomentana. When these have been carefully examined, there will
still remain many interesting monuments, of considerable historical
importance, in other less famous cemeteries; but enough will have
been seen to give an excellent general acquaintance with the main
characteristics of Roma Sotterranea.
PRINTED BY BALLANTYNE, HANSON AND CO.

EDINBURGH AND LONDON
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Chapter 01: An Overview of Evolutionary Biology

1. Charles Darwin’s theory of evolution resulted in a because it changed the way we

5. What is evolutionary biology?

8. Which process is demonstrated in the figure shown?

a. artificial selection c. genetics

9. In what way are natural selection and artificial selection similar?

10. In what way is artificial selection different from natural selection?

11. Which of the following is an example of artificial selection?

13. Antibiotic resistance evolves quickly because

14. Why do physicians often prescribe antibiotics in combination?

22. Which of the following is an example of observational research?

23. Manipulative experiments

29. Mathematical models are useful for

DIF: Easy REF: 1.0

2. Describe Charles Darwin’s theory of evolution by natural selection.

DIF: Easy REF: 1.0

DIF: Moderate REF: 1.0

DIF: Easy REF: 1.1

DIF: Moderate REF: 1.1

DIF: Moderate REF: 1.1

DIF: Moderate REF: 1.1

DIF: Moderate REF: 1.1

DIF: Easy REF: 1.2

DIF: Easy REF: 1.2

DIF: Moderate REF: 1.2

DIF: Difficult REF: 1.2

DIF: Difficult REF: 1.2

DIF: Difficult REF: 1.2

“Heraclius forbad those who had fallen away [in times of

Having sufficiently considered the form of the inscription, let us

We have not promised to conduct the visitor to everything that is

ASPICE, DESCENSU EXSTRUCTO TENEBRISQUE FUGATIS,

“Behold, a new staircase having been made, and the darkness

The second inscription De Rossi would restore as follows:—

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